Habitat conditions and not moss composition mediate microbial community structure in Swiss peatlands.

Peatlands, one of the oldest ecosystems, globally store significant amounts of carbon and freshwater. However, they are under severe threat from human activities, leading to changes in water, nutrient and temperature regimes in these delicate systems. Such shifts can trigger a substantial carbon flux into the atmosphere and diminish the water-holding capacity of peatlands. Microbes associated with moss in peatlands play a crucial role in providing these ecosystem services, which are at risk due to global change. Therefore, understanding the factors influencing microbial composition and function is vital. Our study focused on five peatlands along an altitudinal gradient in Switzerland, where we sampled moss on hummocks containing Sarracenia purpurea. Structural equation modelling revealed that habitat condition was the primary predictor of community structure and directly influenced other environmental variables. Interestingly, the microbial composition was not linked to the local moss species identity. Instead, microbial communities varied significantly between sites due to differences in acidity levels and nitrogen availability. This finding was also mirrored in a co-occurrence network analysis, which displayed a distinct distribution of indicator species for acidity and nitrogen availability. Therefore, peatland conservation should take into account the critical habitat characteristics of moss-associated microbial communities.

Shrinking Alpine chamois: higher spring temperatures over the last 27 years in Switzerland are linked to a 3 kg reduction in body mass of yearlings.

Although climate change is considered to be partly responsible for the size change observed in numerous species, the relevance of this hypothesis for ungulates remains debated. We used body mass measurements of 5635 yearlings (i.e. 1.5 years old) of Alpine chamois (Rupicapra rupicapra) harvested in September in the Swiss Alps (Ticino canton) from 1992 to 2018. In our study area, during this period, yearlings shrank by ca 3 kg while temperatures between May and July rose by 1.7°C. We identified that warmer temperatures during birth and the early suckling period (9 May to 2 July in the year of birth) had the strongest impact on yearling mass. Further analyses of year-detrended mass and temperature data indicate that this result was not simply due to changes in both variables over years, but that increases in temperature during this particularly sensitive time window for development and growth are responsible for the decrease in body mass of yearling chamois. Altogether, our results suggest that rising temperatures in the Alpine regions could significantly affect the ecology and evolution of this wild ungulate.

Theoretical investigation of the dependence of population parameters on initial population size.

In a series of experiments with yeast, classical dynamical models were fitted to populations that differed only in their initial population size (Pylvänäinen 2005). The results revealed a surprising dependence between estimated growth rate and initial population size. Perceived as an artefact, this undesired relationship was tentatively removed by an ad-hoc procedure. This strategy reflects the usual approach of population models in which parameters are not considered to depend on initial conditions. However, our analysis reveals that the observed relationship between estimated growth rate and initial population size is unavoidable when the dimension of a system is reduced. For the present case, the two-dimensional food-yeast system was reduced to a model for yeast only. The consequence of system reduction questions our conception of one-dimensional population models.

Trait-Environment Relationships Reveal the Success of Alien Plants Invasiveness in an Urbanized Landscape.

Urban areas are being affected by rapidly increasing human-made pressures that can strongly homogenize biodiversity, reduce habitat heterogeneity, and facilitate the invasion of alien species. One of the key concerns in invaded urban areas is comparing the trait-environment relationships between alien and native species, to determine the underlying causes of invasiveness. In the current study, we used a trait-environment dataset of 130 native plants and 33 alien plants, recorded in 100 plots covering 50 urban areas and 50 non-urban ones in an urbanization gradient in the arid mountainous Saint-Katherine protected area in Egypt. We measured eleven morphological plant traits for each plant species and ten environmental variables in each plot, including soil resources and human-made pressures, to construct trait-environment associations using a fourth-corner analysis. In addition, we measured the mean functional and phylogenetic distances between the two species groups along an urbanization gradient. Our results revealed strongly significant relationships of alien species traits with human-made pressures and soil resources in urban areas. However, in non-urban areas, alien species traits showed weak and non-significant associations with the environment. Simultaneously, native plants showed consistency in their trait-environment relationships in urban and non-urban areas. In line with these results, the functional and phylogenetic distances declined between the aliens and natives in urban areas, indicating biotic homogenization with increasing urbanization, and increased in non-urban areas, indicating greater divergence between the two species groups. Thereby, this study provided evidence that urbanization can reveal the plasticity of alien species and can also be the leading cause of homogenization in an arid urban area. Future urban studies should investigate the potential causes of taxonomic, genetic, and functional homogenization in species composition in formerly more diverse urbanized areas.

Contrasting alien effects on native diversity along biotic and abiotic gradients in an arid protected area.

Alien impact on native diversity could be a function of both the relatedness of alien species to native community and resources availability. Here, we investigated whether alien plants expand or decrease the functional and phylogenetic space of native plant communities, and how this is affected by alien relatedness to natives and by resources availability. We used a trait-environment dataset of 33 alien and 130 native plants in 83 pairs of invaded and non-invaded plots, covering a gradient of soil resources (organic matter-nitrogen) in Saint-Katherine-Protectorate, Egypt. First, we compared the changes in native composition and calculated alien relatedness to natives within each pair of plots. Second, we tested the effects of resources availability and relatedness on the magnitude of alien impact (defined as a change in native diversity). We found that native composition was phylogenetically less but functionally more diverse in invaded plots compared to non-invaded ones. Moreover, in resources-rich plots, dissimilar aliens to natives significantly increased native diversity, while in resource-limited ones, similar aliens to natives declined native diversity. These results suggest that the assessment of alien impacts in arid-regions is significantly linked to resources-availability and relatedness to natives. Hence, future studies should test the generality of our findings in different environments.

The effects of temperature and dispersal on species diversity in natural microbial metacommunities.

Dispersal is key for maintaining biodiversity at local- and regional scales in metacommunities. However, little is known about the combined effects of dispersal and climate change on biodiversity. Theory predicts that alpha-diversity is maximized at intermediate dispersal rates, resulting in a hump-shaped diversity-dispersal relationship. This relationship is predicted to flatten when competition increases. We anticipate that this same flattening will occur with increased temperature because, in the rising part of the temperature performance curve, interspecific competition is predicted to increase. We explored this question using aquatic communities of Sarracenia purpurea from early- and late-successional stages, in which we simulated four levels of dispersal and four temperature scenarios. With increased dispersal, the hump shape was observed consistently in late successional communities, but only in higher temperature treatments in early succession. Increased temperature did not flatten the hump-shape relationship, but decreased the level of alpha- and gamma-diversity. Interestingly, higher temperatures negatively impacted small-bodied species. These metacommunity-level extinctions likely relaxed interspecific competition, which could explain the absence of flattening of the diversity-dispersal relationship. Our findings suggest that climate change will cause extinctions both at local- and global- scales and emphasize the importance of intermediate levels of dispersal as an insurance for local diversity.

Predator traits determine food-web architecture across ecosystems.

Predator-prey interactions in natural ecosystems generate complex food webs that have a simple universal body-size architecture where predators are systematically larger than their prey. Food-web theory shows that the highest predator-prey body-mass ratios found in natural food webs may be especially important because they create weak interactions with slow dynamics that stabilize communities against perturbations and maintain ecosystem functioning. Identifying these vital interactions in real communities typically requires arduous identification of interactions in complex food webs. Here, we overcome this obstacle by developing predator-trait models to predict average body-mass ratios based on a database comprising 290 food webs from freshwater, marine and terrestrial ecosystems across all continents. We analysed how species traits constrain body-size architecture by changing the slope of the predator-prey body-mass scaling. Across ecosystems, we found high body-mass ratios for predator groups with specific trait combinations including (1) small vertebrates and (2) large swimming or flying predators. Including the metabolic and movement types of predators increased the accuracy of predicting which species are engaged in high body-mass ratio interactions. We demonstrate that species traits explain striking patterns in the body-size architecture of natural food webs that underpin the stability and functioning of ecosystems, paving the way for community-level management of the most complex natural ecosystems.

Increased Temperature Disrupts the Biodiversity-Ecosystem Functioning Relationship.

Gaining knowledge of how ecosystems provide essential services to humans is of primary importance, especially with the current threat of climate change. Yet little is known about how increased temperature will impact the biodiversity-ecosystem functioning (BEF) relationship. We tackled this subject theoretically and experimentally. We developed a BEF theory based on mechanistic population dynamic models, which allows the inclusion of the effect of temperature. Using experimentally established relationships between attack rate and temperature, the model predicts that temperature increase will intensify competition, and consequently the BEF relationship will flatten or even become negative. We conducted a laboratory experiment with natural microbial microcosms, and the results were in agreement with the model predictions. The experimental results also revealed that an increase in both temperature average and variation had a more intense effect than an increase in temperature average alone. Our results indicate that under climate change, high diversity may not guarantee high ecosystem functioning.

Understanding negative biodiversity-ecosystem functioning relationship in semi-natural wildflower strips.

Studies on biodiversity-ecosystem functioning (BEF) in highly controlled experiments often yield results incompatible with observations from natural systems: experimental results often reveal positive relationships between diversity and productivity, while for natural systems, zero or even negative relationships have been reported. The discrepancy may arise due to a limited or closed local species pool in experiments, while natural systems in meta-community contexts experience dynamic processes, i.e., colonization and extinctions. In our study, we analysed plant community properties and above-ground biomass within a semi-natural (i.e., not weeded) experiment in an agricultural landscape. Eleven replicates with four different diversity levels were created from a species pool of 20 wildflower species. We found an overall significant negative relationship between total diversity and productivity. This relationship likely resulted from invasion resistance: in plots sown with low species numbers, we observed colonization by low-performing species; colonization increased species richness but did not contribute substantially to productivity. Interestingly, when analysing the biomass of the sown and the colonizer species separately, we observed in both cases positive BEF relationships, while this relationship was negative for the whole system. A structural equation modelling approach revealed that higher biomass of the sown species was linked to higher species richness, while the positive BEF relationship of the colonizers was indirect and constrained by the sown species biomass. Our results suggest that, in semi-natural conditions common in extensive agroecosystems, the negative BEF relationship results from the interplay between local dominant species and colonization from the regional species pool by subordinate species.

The feasibility of equilibria in large ecosystems: A primary but neglected concept in the complexity-stability debate.

The consensus that complexity begets stability in ecosystems was challenged in the seventies, a result recently extended to ecologically-inspired networks. The approaches assume the existence of a feasible equilibrium, i.e. with positive abundances. However, this key assumption has not been tested. We provide analytical results complemented by simulations which show that equilibrium feasibility vanishes in species rich systems. This result leaves us in the uncomfortable situation in which the existence of a feasible equilibrium assumed in local stability criteria is far from granted. We extend our analyses by changing interaction structure and intensity, and find that feasibility and stability is warranted irrespective of species richness with weak interactions. Interestingly, we find that the dynamical behaviour of ecologically inspired architectures is very different and richer than that of unstructured systems. Our results suggest that a general understanding of ecosystem dynamics requires focusing on the interplay between interaction strength and network architecture.

Effects of temperature variability on community structure in a natural microbial food web.

Climate change research has demonstrated that changing temperatures will have an effect on community-level dynamics by altering species survival rates, shifting species distributions, and ultimately, creating mismatches in community interactions. However, most of this work has focused on increasing temperature, and still little is known about how the variation in temperature extremes will affect community dynamics. We used the model aquatic community held within the leaves of the carnivorous plant, Sarracenia purpurea, to test how food web dynamics will be affected by high temperature variation. We tested the community response of the first (bacterial density), second (protist diversity and composition), and third trophic level (predator mortality), and measured community respiration. We collected early and late successional stage inquiline communities from S. purpurea from two North American and two European sites with similar average July temperature. We then created a common garden experiment in which replicates of these communities underwent either high or normal daily temperature variation, with the average temperature equal among treatments. We found an impact of temperature variation on the first two, but not on the third trophic level. For bacteria in the high-variation treatment, density experienced an initial boost in growth but then decreased quickly through time. For protists in the high-variation treatment, alpha-diversity decreased faster than in the normal-variation treatment, beta-diversity increased only in the European sites, and protist community composition tended to diverge more in the late successional stage. The mortality of the predatory mosquito larvae was unaffected by temperature variation. Community respiration was lower in the high-variation treatment, indicating a lower ecosystem functioning. Our results highlight clear impacts of temperature variation. A more mechanistic understanding of the effects that temperature, and especially temperature variation, will have on community dynamics is still greatly needed.

Persist or Produce: A Community Trade-Off Tuned by Species Evenness.

Understanding the effects of biodiversity on community persistence and productivity is key to managing both natural and production systems. Because rare species face greater danger of extinction, species evenness, a measure of how similar abundances are across species in a community, is seen as a key component of biodiversity. However, previous studies have failed to find a consistent association of species evenness with species survival and biomass production. Here we provide a theoretical framework for the relationship among these three elements. We demonstrate that the lack of consistent outcomes is not an idiosyncratic artifact of different studies but can be unified under one common framework. Applying a niche theory approach, we confirm that under demographic stochasticity evenness is a general indicator of the risk of future species extinctions in a community, in accordance with the majority of empirical studies. In contrast, evenness cannot be used as a direct indicator of the level of biomass production in a community. When a single species dominates, as expressed by the constraints imposed by the population dynamics, biomass production depends on the niche position of the dominating species and can increase or decrease with evenness. We demonstrate that high species evenness and an intermediate level of biomass production is the configuration that maximizes the average species survival probability in response to demographic stochasticity.

Mismatch in microbial food webs: predators but not prey perform better in their local biotic and abiotic conditions.

Understanding how trophic levels respond to changes in abiotic and biotic conditions is key for predicting how food webs will react to environmental perturbations. Different trophic levels may respond disproportionately to change, with lower levels more likely to react faster, as they typically consist of smaller-bodied species with higher reproductive rates. This response could cause a mismatch between trophic levels, in which predators and prey will respond differently to changing abiotic or biotic conditions. This mismatch between trophic levels could result in altered top-down and bottom-up control and changes in interaction strength. To determine the possibility of a mismatch, we conducted a reciprocal-transplant experiment involving Sarracenia purpurea food webs consisting of bacterial communities as prey and a subset of six morphologically similar protozoans as predators. We used a factorial design with four temperatures, four bacteria and protozoan biogeographic origins, replicated four times. This design allowed us to determine how predator and prey dynamics were altered by abiotic (temperature) conditions and biotic (predators paired with prey from either their local or non-local biogeographic origin) conditions. We found that prey reached higher densities in warmer temperature regardless of their temperature of origin. Conversely, predators achieved higher densities in the temperature condition and with the prey from their origin. These results confirm that predators perform better in abiotic and biotic conditions of their origin while their prey do not. This mismatch between trophic levels may be especially significant under climate change, potentially disrupting ecosystem functioning by disproportionately affecting top-down and bottom-up control.

Matching-centrality decomposition and the forecasting of new links in networks.

Networks play a prominent role in the study of complex systems of interacting entities in biology, sociology, and economics. Despite this diversity, we demonstrate here that a statistical model decomposing networks into matching and centrality components provides a comprehensive and unifying quantification of their architecture. The matching term quantifies the assortative structure in which node makes links with which other node, whereas the centrality term quantifies the number of links that nodes make. We show, for a diverse set of networks, that this decomposition can provide a tight fit to observed networks. Then we provide three applications. First, we show that the model allows very accurate prediction of missing links in partially known networks. Second, when node characteristics are known, we show how the matching-centrality decomposition can be related to this external information. Consequently, it offers us a simple and versatile tool to explore how node characteristics explain network architecture. Finally, we demonstrate the efficiency and flexibility of the model to forecast the links that a novel node would create if it were to join an existing network.

Top predators affect the composition of naive protist communities, but only in their early-successional stage.

Introduced top predators have the potential to disrupt community dynamics when prey species are naive to predation. The impact of introduced predators may also vary depending on the stage of community development. Early-succession communities are likely to have small-bodied and fast-growing species, but are not necessarily good at defending against predators. In contrast, late-succession communities are typically composed of larger-bodied species that are more predator resistant relative to small-bodied species. Yet, these aspects are greatly neglected in invasion studies. We therefore tested the effect of top predator presence on early- and late-succession communities that were either naive or non-naive to top predators. We used the aquatic community held within the leaves of Sarracenia purpurea. In North America, communities have experienced the S. purpurea top predator and are therefore non-naive. In Europe, this predator is not present and its niche has not been filled, making these communities top-predator naive. We collected early- and late-succession communities from two non-naive and two naive sites, which are climatically similar. We then conducted a common-garden experiment, with and without the presence of the top predator, in which we recorded changes in community composition, body size spectra, bacterial density, and respiration. We found that the top predator had no statistical effect on global measures of community structure and functioning. However, it significantly altered protist composition, but only in naive, early-succession communities, highlighting that the state of community development is important for understanding the impact of invasion.

The importance of landscape and spatial structure for hymenopteran-based food webs in an agro-ecosystem.

1. Understanding the environmental factors that structure biodiversity and food webs among communities is central to assess and mitigate the impact of landscape changes. 2. Wildflower strips are ecological compensation areas established in farmland to increase pollination services and biological control of crop pests and to conserve insect diversity. They are arranged in networks in order to favour high species richness and abundance of the fauna. 3. We describe results from experimental wildflower strips in a fragmented agricultural landscape, comparing the importance of landscape, of spatial arrangement and of vegetation on the diversity and abundance of trap-nesting bees, wasps and their enemies, and the structure of their food webs. 4. The proportion of forest cover close to the wildflower strips and the landscape heterogeneity stood out as the most influential landscape elements, resulting in a more complex trap-nest community with higher abundance and richness of hosts, and with more links between species in the food webs and a higher diversity of interactions. We disentangled the underlying mechanisms for variation in these quantitative food web metrics. 5. We conclude that in order to increase the diversity and abundance of pollinators and biological control agents and to favour a potentially stable community of cavity-nesting hymenoptera in wildflower strips, more investment is needed in the conservation and establishment of forest habitats within agro-ecosystems, as a reservoir of beneficial insect populations.

Diversity protects plant communities against generalist molluscan herbivores.

Wildflower strips are used to increase natural enemies of crop pests and to conserve insect diversity on farmland. Mollusks, especially slugs, can affect the vegetation development in these strips considerably. Although recent theoretical work suggests that more diverse plant communities will exhibit greater resistance against herbivore pressure, empirical studies are scarce. We conducted a semi-natural experiment in wildflower strips, manipulating trophic structure (reduction in herbivorous mollusks and reduction in major predators) and plant diversity (2, 6, 12, 20 and 24 sown species). This design allowed us to assess the effect of plant diversity, biomass and composition on mollusks, and vice versa, the effect of mollusc abundance on vegetation. Seven species of mollusks were found in the strips, with the slugs Arion lusitanicus, Deroceras reticulatum and Deroceras panormitanum being most frequent. We found a negative relationship between plant diversity and mollusk abundance, which was due predominantly to a decrease in the agricultural pest species A. lusitanicus. These results are consistent with the hypothesis that plant diversity can reduce the impact of herbivores. However, plant identity also had an effect on mollusks, and accounted for a much larger fraction of the variation in mollusk communities than biodiversity effects. While overall plant diversity decreased during the 3 years of the study, in the final year the highest plant diversity was found in the plots where mollusk populations were experimentally reduced. We conclude that selective feeding by generalist herbivores leads to changes in plant community composition and hence reduced plant diversity. Our results highlight the importance of plant biodiversity as protection against generalist herbivores, which if abundant can in the long term negatively impact plant diversity, driving the system along a "low plant diversity - high mollusk abundance" trajectory.

Phylogeny versus body size as determinants of food web structure.

Food webs are the complex networks of trophic interactions that stoke the metabolic fires of life. To understand what structures these interactions in natural communities, ecologists have developed simple models to capture their main architectural features. However, apparently realistic food webs can be generated by models invoking either predator-prey body-size hierarchies or evolutionary constraints as structuring mechanisms. As a result, this approach has not conclusively revealed which factors are the most important. Here we cut to the heart of this debate by directly comparing the influence of phylogeny and body size on food web architecture. Using data from 13 food webs compiled by direct observation, we confirm the importance of both factors. Nevertheless, phylogeny dominates in most networks. Moreover, path analysis reveals that the size-independent direct effect of phylogeny on trophic structure typically outweighs the indirect effect that could be captured by considering body size alone. Furthermore, the phylogenetic signal is asymmetric: closely related species overlap in their set of consumers far more than in their set of resources. This is at odds with several food web models, which take only the view-point of consumers when assigning interactions. The echo of evolutionary history clearly resonates through current food webs, with implications for our theoretical models and conservation priorities.

Ecophylogenetics: advances and perspectives.

Ecophylogenetics can be viewed as an emerging fusion of ecology, biogeography and macroevolution. This new and fast-growing field is promoting the incorporation of evolution and historical contingencies into the ecological research agenda through the widespread use of phylogenetic data. Including phylogeny into ecological thinking represents an opportunity for biologists from different fields to collaborate and has provided promising avenues of research in both theoretical and empirical ecology, towards a better understanding of the assembly of communities, the functioning of ecosystems and their responses to environmental changes. The time is ripe to assess critically the extent to which the integration of phylogeny into these different fields of ecology has delivered on its promise. Here we review how phylogenetic information has been used to identify better the key components of species interactions with their biotic and abiotic environments, to determine the relationships between diversity and ecosystem functioning and ultimately to establish good management practices to protect overall biodiversity in the face of global change. We evaluate the relevance of information provided by phylogenies to ecologists, highlighting current potential weaknesses and needs for future developments. We suggest that despite the strong progress that has been made, a consistent unified framework is still missing to link local ecological dynamics to macroevolution. This is a necessary step in order to interpret observed phylogenetic patterns in a wider ecological context. Beyond the fundamental question of how evolutionary history contributes to shape communities, ecophylogenetics will help ecology to become a better integrative and predictive science.

Phylogenetic signal in predator-prey body-size relationships.

Body mass is a fundamental characteristic that affects metabolism, life history, and population abundance and frequently sets bounds on who eats whom in food webs. Based on a collection of topological food webs, Ulrich Brose and colleagues presented a general relationship between the body mass of predators and their prey and analyzed how mean predator-prey body-mass ratios differed among habitats and predator metabolic categories. Here we show that the general body-mass relationship conceals significant variation associated with both predator and prey phylogeny. Major-axis regressions between the log body mass of predators and prey differed among taxonomic groups. The global pattern for Kingdom Animalia had slope > 1, but phyla and classes varied, and several had slopes significantly < 1. The predator-prey body-mass ratio can therefore decrease or increase with increasing body mass, depending on the taxon considered. We also found a significant phylogenetic signal in analyses of prey body-mass range for predators and predator body-mass range for prey, with stronger signal in the former. Besides providing insights into how characteristics of trophic interactions evolve, our results emphasize the need to integrate phylogeny to improve models of community structure and dynamics or to achieve a metabolic theory of food-web ecology.